Territory quality affects female preference in a Lake Victoria cichlid fish

Mate preferences on male colour have been implicated in generating and maintaining species diversity among haplochromine cichlid fish. Their lek-like mating system suggests that not only male colour but also territory quality is instrumental in mate choice. We assessed the relative importance of territory quality and male colour in mate choice by testing whether territory quality can override the female preference for males of her own colour in the Lake Victoria cichlid genus Pundamilia. First, we showed in experimental groups that the dominant male preferentially monopolised a large tube relative to a small tube. The situation mimics quality difference in rocky crevices that serves as a focal point for male courtship display. Second, in mate-choice tests, Pundamilia nyererei females were allowed to choose between closely related P. nyererei and P. pundamilia males; these species differ strikingly in male nuptial coloration, but little else. We gave either both males the same small tube or one of them a large tube. The preference of P. nyererei females for P. nyererei males in the control situation (where both males had a small tube) was significantly diminished in favour of P. pundamilia males when the latter had the large tube. The results provide experimental evidence that differences in territory quality can override the female preference for males of her own colour. This finding is critical for a recent hypothesis proposing that male competition for mating territories can facilitate the process of sympatric speciation by sexual selection.     

Immune function, dominance and mating success in drumming male wolf spiders Hygrolycosa rubrofasciata

In this study, we provide a piece of experimental evidence that immune function is related to dominance and mating success in wild caught male wolf spiders, Hygrolycosa rubrofasciata. In the mating season, H. rubrofasciata males are actively searching for receptive females, and while searching males often engage in agonistic behavior (i.e., agonistic drumming signals, chases, and fights) with each other. The present results demonstrate that dominant males had higher lytic activities in their hemolymph than subordinates. Lytic activity estimates the concentration of antimicrobial peptides with lysozyme-like activity in hemolymph, which have been shown to play an important role in defense against bacteria, viruses, and fungi. Dominants also had higher courtship drumming rates than subordinates. Moreover, winners in mating competition had higher lytic activities than losers, but this was measured nonindependently of dominance status. Among males with mating failure, there was a moderate negative correlation between encapsulation rate and courtship drumming rate, suggesting that low quality males might not be able to bear the immunological costs of courtship behavior. These results suggest that females might gain immunological benefits by preferring vigorous males.     

Male signaling behavior and sexual selection in a wolf spider (Araneae: Lycosidae): a test for dual functions

Male signaling behaviors are often studied in a single context but may serve multiple functions (e.g., in male–male competition and female mate choice). We examined the issue of dual function male signals in a wolf spider species Schizocosa ocreata (Hentz) that displays the same species-specific signaling behaviors in both male–male and male–female contexts. These signaling behaviors have been described as either aggression or courtship according to the context observed. We tested the possibility of dual functions by comparing the relationship between behaviors and outcome of male–male contests (winner/loser) and male–female mating encounters (mating success). Frequency, rate, and mean duration of signaling behaviors did not vary with outcome of male–male contests, which appears instead to be based upon relative size and body mass. Winners of contests had significantly greater body mass than losers, and greater mass relative to opponents was significantly associated with probability of winning. Overall, signaling rates were much higher in male–female interactions than in male–male contests and were higher for males that successfully mated than for those that did not mate. Mean duration of some male displays was also greater for males that successfully mated. However, male size was not associated with probability of mating. Taken together, results suggest an intersexual selection context for the current function of male signals in these wolf spiders and that increased display vigor is associated with male mating success.     

Can male-male competition stabilize speciation? A test in Lake Victoria haplochromine cichlid fish

It has been suggested that sympatric speciation can be driven by sexual selection on male mating traits alone. However, a fundamental problem for this process is the lack of ecological differentiation that would stabilize the coexistence of incipient species through frequency-dependent selection. Such selection can also occur if male aggression is primarily directed towards similar rather than towards dissimilar phenotypes, so that rare male phenotypes would enjoy a negatively frequency-dependent fitness advantage. We experimentally tested such an aggression bias in two recently diverged, ecologically and anatomically similar sympatric cichlid species pairs of the genus Pundamilia from Lake Victoria. Territorial males of a pair of partially reproductively isolated species with red and blue nuptial coloration, respectively, studied in the laboratory were confronted simultaneously with both colour types enclosed in transparent tubes. Red males were more aggressive to red stimuli under white light but not when colour differences were masked under green light. Blue males were equally aggressive to both stimuli in both light conditions. Males of two apparently fully reproductively isolated species, again one with red and one with blue nuptial coloration, studied in the field, both directed more aggressive behaviour towards conspecific than towards heterospecific stimulus males. The differential allocation of aggression would create an advantage for males of the less abundant phenotype or species, thereby potentially supporting stable coexistence of the phenotypes. The finding that this effect was less clear in the partially reproductively isolated species pair than in the fully isolated species pair is discussed.     

Determinants of male territorial behavior in a Hungarian collared flycatcher population: plumage traits of residents and challengers

In intrasexual conflicts, contestants can rely on relative or absolute size of status badges to mediate aggressive behavior. Most studies focus on the response of focal animals to variation in status badges of their competitors; few have simultaneously considered the traits of both participants under experimental conditions. By simulating territorial intrusions, we tested the importance of two sexual traits [forehead patch size (FPS) and wing patch size (WPS)] in territorial behavior of males in a Hungarian population of the collared flycatcher, Ficedula albicollis. We presented a stimulus male to an unpaired resident male to examine whether the characteristics of the territory owner or those of the challenger were associated with the latency of the first attack of the owner, which is a good predictor of the territorial behavior in general. WPS of the stimulus male was a significant determinant of the latency of the first attack, as males with a larger WPS elicited quicker attacks from the residents than males with a small WPS. From the residents’ perspective, age appeared to influence their territorial behavior, as yearlings had shorter attack latencies than older males. Additionally, latency could be considered an individual-specific attribute because it varied consistently among males, even when the WPS of the stimulus male was controlled, and it was associated with pairing success. Contrary to findings in a Swedish population, FPS seemed to be unimportant in male–male competition in our population, which suggests population differences in the role of the two plumage traits. Our results indicate that in a territorial conflict, the characteristics of both participants are important.     

Armaments and ornaments in the rock sparrow: a possible dual utility of a carotenoid-based feather signal

Females often base their mating preferences on male sexual secondary traits that are used to settle contests among males. Such traits are likely to be honest indicators of male quality if they are constantly used during costly male–male agonistic interactions. Carotenoid signals have been shown to work as a handicap because they are costly to produce. However, the role of carotenoids as “honest” signals during male contests is less clear, and it is not known whether a carotenoid-based trait can serve in both male–male competition and female choice. In this study, we studied the dual function of a carotenoid feather ornament in the rock sparrow (Petronia petronia), a bird species in which both sexes have a yellow throat patch whose size positively correlates with phenotypic measures. First, we investigated, in a field study, whether the size of a male’s yellow patch correlates with his ability to acquire a territory. Second, we tested the signal function of the yellow patch in two male–male interaction in captivity experiments. Finally, we measured female preference for males differing in throat patch size in a mate choice experiment. Our experiments revealed that the size of a male’s throat patch positively correlated with the number of nest boxes he was able to defend. Moreover, in controlled conditions, males with relatively large yellow patches had earlier access to food than those with small patches. Also, in an experiment in which a dummy rock sparrow with an experimentally manipulated yellow patch was positioned near a feeder, latency to feed by focal birds positively correlated with dummy patch size. Lastly, in a dichotomous mate choice experiment, females showed a proximity preference for males whose patch was experimentally enlarged. Taken together, these results suggest that the same carotenoid feather signal may be used in both male–male competition and female choice in this passerine bird.     

Vocal interactions in common nightingales (<Emphasis Type="Italic">Luscinia megarhynchos</Emphasis>): males take it easy after pairing

Seasonal patterns of bird song have been studied intensively with a focus on individual males. However, little is known about seasonal patterns of singing during vocal interactions between males. Vocal interactions have been shown to be important in sexual selection as males may signal aspects of motivation or quality. Here, we investigated in nightingales (Luscinia megarhynchos) whether a male’s behaviour in vocal interactions at different stages of the breeding season is influenced by its mating status. We examined how males that differ in their subsequent mating success respond to a non-interactive, nocturnal playback presented during the period of mate attraction and subsequently during the egg-laying period. We found that mated males overlapped fewer songs and had a lower song rate during the egg-laying period compared to their responses during the mate-attraction period, whereas unpaired males did not vary in their responses between the two periods. Our results suggest that mating status is a key factor affecting singing behaviour in vocal interactions and that a time-specific singing pattern like song overlapping is used flexibly during vocal interactions. Because song overlapping is thought to be a signal of aggression in male–male vocal interactions, it seems that males vary the level of aggression in vocal interactions according to their mating status and to the stage in the breeding season.     

Ectoparasites of house sparrows (Passer domesticus): an experimental test of the immunocompetence handicap hypothesis and a new model

Conspicuous secondary sexual traits may have evolved as handicap-revealing signals or as badges of status. We present results of an experiment using males of the sexually dimorphic house sparrow (Passer domesticus), that support the idea that the male-specific bib can be both a handicap-revealing signal and a reliable badge indicating the physical condition of the bird. In a test of the immunocompetence handicap hypothesis, wild-caught adult male house sparrows were studied in captivity. Birds implanted with elevated doses of testosterone were more dominant, had higher circulating levels of both testosterone and corticosterone and they also harboured relatively larger ectoparasite loads. Higher parasite loads were also associated with individuals showing lower immunocompetence and larger changes in bib size. A new model for immunocompetence effects in sexual selection is introduced, integrating actions that the hypothalamopituitary axis exerts on gonads, adrenals and the thyroid gland. The ”integrated immunocompetence model” synthesizes both the ”handicap” (i.e. survival-decreasing) and ”badge of status” (i.e. survival- enhancing) models for evolution of secondary sexual traits. Received: 15 May 1999 / Received in revised form: 2 November 1999 / Accepted: 6 November 1999     

Advertisement-call modification,male competition,and female preference in the bird-voiced treefrog <Emphasis Type="Italic">Hyla avivoca</Emphasis>

Senders and receivers influence dynamic characteristics of the signals used for mate attraction over different time scales. On a moment-to-moment basis, interactions among senders competing for a mate influence dynamic characteristics, whereas the preferences of receivers of the opposite gender exert an influence over evolutionary time. We observed and recorded the calling patterns of the bird-voiced treefrog Hyla avivoca to assess how the dynamic characters of calls vary during interactions among groups of males in a chorus. This question was also addressed using playback experiments with males. Playback experiments with females showed how changes in dynamic call properties are likely to affect male mating success. Frogs calling in pairs, groups, or in response to playbacks produced longer calls than did isolated males. During call overlap, males often increased the duration of the silent interval (gaps) between the pulses of their calls so that the pulses of the calls of two neighbors interdigitated. This change resulted in increased variability of pulse rate, a traditionally static acoustic property; however, males also produced high proportions of non-overlapped calls in which variability in pulse rate was low and had species-typical values. Females preferred long calls to short- and average-duration calls, and non-overlapped calls to overlapped calls. Given a choice between pairs of overlapped calls, females preferred pairs in which the proportion of overlap was low and pairs in which the pulses of such calls interdigitated completely. The observed patterns of vocal competition thus reflect the preferences of conspecific females, which have influenced the evolution of the calling behavior of H. avivoca. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Maintenance of variance in sperm transfer rates in a scorpionfly: food availability,genetic basis,and heritability

Variation in traits closely related to fitness is expected to be low. This is because these traits are under directional selection and the best genotype should prevail. However, there have been a number of studies demonstrating the existence of considerable variance in sexually selected traits, which is generally known as the lek paradox. Accordingly, earlier studies found substantial variation in sperm transfer rates in Panorpa vulgaris. Aiming at finding the mechanism that maintains this variation, we analyzed the condition dependence and the narrow sense heritability of sperm transfer rates. Food deprivation in the larval/adult phase caused a decrease in the males’ capability of saliva secretion resulting in shorter copulations and a reduced number of transferred sperm. There was a positive correlation between mean sperm transfer rates and mean body mass. Additionally, intermale variation in sperm transfer rates decreased with increasing food availability. Hence, we suggest that sperm transfer rates in P. vulgaris are influenced by adult feeding history. Heritability analyses of sperm transfer rates did not provide significant results, which is consistent with the general hypothesis that additive genetic variance in traits closely related to fitness is small. Since a trait’s potential to respond to selection is proportional to the amount of contained additive genetic variance, the ascertained small heritability provides a satisfying explanation for the maintenance of substantial variation in sperm transfer rates.     

Early nutritional conditions, growth trajectories and mate choice: does compensatory growth lead to a reduction in adult sexual attractiveness?

It is well documented that poor early nutrition can have profound negative effects on adult life-history traits. However, it has also been demonstrated that organisms can undergo compensatory resource allocation strategies (such as an accelerated growth rate) if food availability improves, so as to mitigate the effects of the poor early conditions. Previous research has indicated that elevated growth rates can incur costs in the longer term, such as an increased rate of senescence and shorter lifespan. We tested whether a phase of compensatory growth after a period of reduced food availability earlier in life affected the sexual attractiveness of adult male green swordtails Xiphophorus helleri, a species in which it has previously been documented that females prefer larger-bodied and longer-tailed males. The experiment compared the attractiveness of size-matched brothers that had experienced contrasting growth trajectories as juveniles; the experiments were initially conducted in the middle of a male’s sexually reproductive life and were then repeated towards the very end of life. At both ages, males that had undergone compensatory growth were equally as attractive as their brothers that had grown normally. These results suggest that the growth compensation benefits males through an increase in their attractiveness over that which they would have had if they had remained on their original growth trajectory. The lack of change in relative attractiveness with age indicates that the compensation does not cause greater deterioration in secondary sexual characters at older ages than in continuously well-fed males.     

Parental investment, adult sex ratios, and sexual selection in a socially monogamous seabird

Although most birds are monogamous, theory predicts that greater female parental investment and female-biased adult sex ratios will lower the polygyny threshold. This should result in polygynous mating, unless obligate biparental care or the spatial and temporal distribution of fertilizable females constrains a male’s ability to take advantage of a lowered polygyny threshold. Here we present data on the extent of male sexually dimorphic plumage, adult sex ratios and breeding season synchrony in three populations of a socially monogamous seabird, the brown booby Sula leucogaster. For one of these populations, San Pedro Mártir Island, we also present data on differences in male and female parental investment, mortality and probability of pairing. The extent of plumage dimorphism varied among populations. Sex ratios were female biased in all populations. On San Pedro Mártir Island, parental investment was female biased, females failed more often than males to find a mate, but there was no polygyny. We suggest that on San Pedro Mártir: (1) a period of obligate biparental care coupled with a relatively synchronous breeding season constrained the ability of males to take advantage of a high environmental polygamy potential and (2) the resulting socially monogamous mating system, in combination with the female-biased adult sex ratio, caused females to be limited by the availability of males despite their greater parental investment. Received: 18 November 1999 / Accepted: 24 January 2000     

Black spots and female association preferences in a sexual/asexual mating complex (Poecilia, Poeciliidae, Teleostei)

We investigated whether female association preferences for males are influenced by black spot disease (BSD), a parasite induced change of the host phenotype. We compared three different species of fish: a gynogenetic hybrid species, Poecilia formosa (amazon molly) and two sexual species (Poecilia latipinna and Poecilia mexicana), which were involved in the natural hybridisation leading to the amazon molly. Contrary to their sexual relatives, asexual amazon mollies significantly avoided images of males infected with black spot disease. We propose that amazon molly females have direct fitness benefits from choosing healthy males. The adaptive significance of the preference for BSD-uninfected males in the asexual amazon molly is yet unclear but may involve avoidance of predation or parasite infection as well as increased sperm availability.